Plant Cell Biology
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Plant Cell Biology 01_P374233_PRELIMS.indd i 8/11/2009 2:35:28 PM 01_P374233_PRELIMS.indd ii 8/11/2009 2:35:28 PM Plant Cell Biology From Astronomy to Zoology Randy Wayne AMSTERDAM • BOSTON • HEIDELBERG • LONDON • NEW YORK • OXFORD PARIS • SAN DIEGO • SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier 01_P374233_PRELIMS.indd iii 8/11/2009 2:35:28 PM Academic Press is an imprint of Elsevier 30 Corporate Drive, Suite 400, Burlington, MA 01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Theobald’s Road, London WC1X 8RR, UK © 2009 Elsevier, Inc. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Details on how to seek permission, further information about the Publisher’s permissions policies and our arrangements with organizations such as the Copyright Clearance Center and the Copyright Licensing Agency, can be found at our website: www.elsevier.com/permissions. This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted herein). Notices Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary. Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the safety of others, including parties for whom they have a professional responsibility. To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideas contained in the material herein. Library of Congress Cataloging-in-Publication Data Wayne, Randy. Plant cell biology / Randy Wayne. p. cm. Includes bibliographical references and index. ISBN 978-0-12-374233-9 (hardback : alk. paper) 1. Plants — Cytology. I. Title. QK725.W39 2009 571.6’2 — dc22 2009018976 British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library. ISBN : 978-0-12-374233-9 For information on all Academic Press publications visit our website at www.elsevierdirect.com Companion Website Available http://www.elsevierdirect.com/companions/9780123742339 09 10 11 12 13 5 4 3 2 1 Printed in the United States of America 02_P374233_ITR.indd iv 8/6/2009 6:18:50 PM Dedicated to President John F. Kennedy for inspiring my generation to be courageous in the pursuit of science 03_P374233_DED.indd v 8/6/2009 6:19:28 PM 03_P374233_DED.indd vi 8/6/2009 6:19:28 PM Contents Preface xiii 2 .12 The Cytoskeleton – Plasma Membrane – Extracellular Matrix 1 . On the Nature of Cells Continuum 49 1 .1 Introduction: What Is a Cell? 1 2 .13 Summary 50 1 .2 The Basic Unit of Life 4 2 .14 Questions 50 1 .3 The Chemical Composition of Cells 6 3. Plasmodesmata 1 .4 A Sense of Cellular Scale 7 3 .1 The Relationship between Cells and 1 .5 The Energetics of Cells 9 the Organism 51 1 .6 Are There Limits to the Mechanistic 3 .2 Discovery and Occurrence of View? 10 Plasmodesmata 52 1 .7 The Mechanistic Viewpoint 3 .3 Structure of Plasmodesmata 53 and God 11 3 .4 Isolation and Composition of 1 .8 What Is Cell Biology? 12 Plasmodesmata 55 1 .9 Summary 12 3 .5 Permeability of Plasmodesmata 56 1 .10 Questions 14 3 .6 Summary 60 3 .7 Questions 60 2 . Plasma Membrane 2 .1 The Cell Boundary 15 4 . Endoplasmic Reticulum 2 .2 Topology of the Cell 15 4 .1 Significance and Evolution of the 2 .3 Evidence for the Existence of a Plasma Endoplasmic Reticulum 61 Membrane 16 4 .2 Discovery of the Endoplasmic 2 .4 Structure of the Plasma Membrane 20 Reticulum 61 2 .5 Isolation of the Plasma Membrane 23 4 .3 Structure of the Endoplasmic 2 .6 Chemical Composition of the Plasma Reticulum 62 Membrane 25 4 .4 Structural Specializations That Relate 2 .7 Transport Physiology 31 to Function 64 2 .8 Electrical Properties of the Plasma 4 .5 Isolation of RER and SER 65 Membrane 39 4 .6 Composition of the ER 65 2 .9 Characterization of Two Transport 4 .7 Function of the Endoplasmic Proteins of the Plasma Membrane 41 Reticulum 65 2.9.1 Proton-Pumping ATPase 41 4.7.1 Lipid Synthesis 65 2.9.2 The K ϩ Channel 44 4.7.2 Protein Synthesis on the 2 .10 Plasma Membrane – Localized Endoplasmic Reticulum 66 Physiological Responses 47 4.7.3 Protein Glycosylation 2.10.1 Guard Cells 47 (Carbohydrate Synthesis) 71 2.10.2 Motor Organs 48 4.7.4 Calcium Regulation 71 2.10.3 Action Potentials 48 4.7.5 Phenylpropanoid and Flavonoid 2.10.4 Cell Polarization 48 Synthesis 73 2 .11 Structural Specializations of 4 .8 Summary 74 the Plasma Membrane 48 4 .9 Questions 74 vii 04_P374233_Prelims1.indd vii 8/13/2009 8:20:44 PM viii Contents 5. Peroxisomes 8. Movement within the Endomembrane 5 .1 Discovery of Microbodies 75 System 5 .2 Isolation of Peroxisomes 75 8 .1 Discovery of the Secretory Pathway 119 5 .3 Composition of Peroxisomes 76 8 .2 Movement to the Plasma Membrane 5 .4 Function of Peroxisomes 76 and the Extracellular Matrix 123 5.4.1 β -Oxidation 76 8.2.1 Movement between the ER 5.4.2 Photorespiration 78 and the Golgi Apparatus 124 5 .5 Relationship between 8.2.2 Movement from the Golgi Glyoxysomes and Peroxisomes 81 Apparatus to the Plasma 5 .6 Metabolite Channeling 83 Membrane 125 5 .7 Other Functions 85 8 .3 Movement from the ER to the Golgi 5 .8 Biogenesis of Peroxisomes 86 Apparatus to the Vacuole 126 5 .9 Evolution of Peroxisomes 88 8 .4 Movement from the ER to the Vacuole 126 5 .10 Summary 88 8 .5 Movement from the Plasma 5 .11 Questions 88 Membrane to the Endomembranes 127 8.5.1 Fluid-Phase Endocytosis 127 6 . Golgi Apparatus 8.5.2 Receptor-Mediated Endocytosis 130 6 .1 Discovery and Structure of the 8.5.3 Piggyback Endocytosis 131 Golgi Apparatus 89 8 .6 Disruption of Intracellular Secretory 6 .2 Polarity of the Golgi Stack 91 and Endocytotic Pathways 131 6 .3 Isolation of the Golgi Apparatus 92 8 .7 Summary 132 6 .4 Composition of the Golgi Apparatus 93 8 .8 Questions 132 6 .5 Function of the Golgi Apparatus 93 6.5.1 Processing of Glycoproteins 93 9. Cytoplasmic Structure 6.5.2 Synthesis of Carbohydrates 93 9 .1 Historical Survey of the Study of 6.5.3 Transport of Sugars 94 Cytoplasmic Structure 133 6 .6 The Mechanism of Movement from 9 .2 Chemical Composition of Cisterna to Cisterna 94 Protoplasm 136 6 .7 Positioning of the Golgi Apparatus 98 9 .3 Physical Properties of Cytoplasm 137 6 .8 Summary 99 9.3.1 Viscosity of the Cytoplasm 138 6 .9 Questions 99 9.3.2 Elasticity of the Cytoplasm 147 9 .4 Microtrabecular Lattice 148 7. The Vacuole 9.4.1 Function of the Microtrabecular 7 .1 Discovery of the Vacuole 101 Lattice in Polarity 148 7 .2 Structure, Biogenesis, and Dynamic 9 .5 Summary 149 Aspects of Vacuoles 102 9 .6 Questions 149 7 .3 Isolation of Vacuoles 105 7 .4 Composition of Vacuoles 106 10. Actin and Microfi lament-Mediated 7 .5 Transport across the Vacuolar Processes Membrane 108 10 .1 Discovery of Actomyosin and the 7.5.1 Proton-Translocating Pumps 109 Mechanism of Muscle Movement 151 7.5.2 ABC (ATP-Binding Cassette) 10 .2 Actin in Nonmuscle Cells 154 Transporters or Traffic ATPases 110 10.2.1 Temporal and Spatial 7.5.3 Slowly Activated Vacuolar Localization of Actin in Channels 110 Plant Cells 154 7.5.4 Water Channels 111 10.2.2 Biochemistry of Actin 155 7 .6 Functions of the Vacuole 111 10.2.3 Biochemistry of Myosins 157 7.6.1 Proteolysis and Recycling 111 10 .3 Force-Generating Reactions Involving 7.6.2 Taking up Space 112 Actin 157 7.6.3 Storage and Homeostasis 113 10.3.1 Actomyosin 157 7.6.4 Role in Turgor Generation 115 10.3.2 Polymerization of Actin 7.6.5 Other Functions 117 Filaments 158 7 .7 Biotechnology 117 10 .4 Actin-Based Motility 158 7 .8 Summary 117 10.4.1 Cytoplasmic Streaming 159 7 .9 Questions 118 10.4.2 Chloroplast Movements 162 04_P374233_Prelims1.indd viii 8/13/2009 8:20:45 PM Contents ix 10.4.3 Cell Plate Reorientation in 12 .5 A Kinetic Description of Regulation 189 Allium 163 12.5.1 Early History of Kinetic 10.4.4 Secretion of Vesicles Involved Studies 189 in Tip Growth and Auxin-Induced 12.5.2 Kinetics of Enzyme Reactions 190 Growth 163 12.5.3 Kinetics of Diffusion and 10.4.5 Contractile Vacuoles 164 Dehydration 194 10 .5 Role of Actin in Membrane Transport 164 12.5.4 A Thermodynamic Analysis of 10 .6 Summary 164 the Signal-to-Noise Problem 196 10 .7 Questions 164 12 .6 Ca 2 ϩ Signaling System 196 12 .7 Mechanics of Doing Experiments to 11 .